Ellescus bipunctatus (Linnaeus, 1758)

Figs. 3A, 4, 5B, 6B

Curculio bipunctatus Linnaeus, 1758 (original description); Paykull 1792: 58 (description, comment).

Curculio unipunctatus Olivier, 1791 (description).

Orchestes ruficornis Zetterstedt, 1837 (description).

Mecinus erythrocerus Abeille de Perrin, 1910 (description).

Elleschus bipunctatus; Hubbard et al. 1878: 621 (taxonomic comment); Austin 1880: 49 (catalogue); Hamilton 1889: 156 (distribution); Slosson 1897: 239 (checklist); Blatchley and Leng 1916: 283 (identification, distribution); Hoffman 1954: 1131 (identification); O’ Brien and Wibmer 1982: 116 (catalogue, distribution); Mattson et al. 1994: 10 (catalogue).

Elleschus bipustulatus; Hatch 1971: 342 (error; catalogue).

Elleschus borealis Carr, 1920 (description); Bousquet et al. 2013: 326 (catalogue, distribution); new synonym.

Ellescus bipunctatus; Bousquet et al. 2013: 326 (catalogue, distribution); Webster et al. 2016: 373–374 (distribution); Alonso-Zarazaga et al. 2017: 196 (catalogue, distribution).

Notes on the type series. This species was described from an unknown number of specimens collected in “Europa”. Type specimens of this species are not present in the Linnaean Society of London collections or the University of Uppsala’ s Museum of Evolution collection and are therefore presumed lost. Although all specimens of E. bipunctatus examined in European collections were accurately identified and the “consensus identity” of this species is relatively stable in Europe, North American specimens of this species frequently have been confused with E. scanicus (see Fig. 4A, B), supporting the designation of a neotype for this species. Here, we designate a male ZSM specimen as the neotype to properly stabilise the identity of this species. We chose the ZSM specimen (Germany) because the original specimens used to describe the species were European in origin and dark in colour. The neotype specimen has, attached, a rectangular red label reading “ NEOTYPE ” with the species name, species author, and year of description, and a line reading “designated Lewis and Anderson” along with the year of designation.

Specimens examined

Neotype (designated here). GERMANY: Bavaria, Holzkirchen, 8.v.1959, K. Gaigl (male, ZSM).

Other material. AUSTRIA: Lower Austria, Tullnerbach, A. Winkler (2, CMNC); Wendbach (1, MIZ), COL000400; CANADA: Alberta: Calgary, 14.vi.1953 – 18.vi.1959, B. and J. Carr (8, CNCI) ; Ghost Dam, 18.vi.1959 – 22.v.1981, B. and J. Carr (7, CNCI) ; Edmonton, vi.1915– 26.iv.1922, F.S. Carr (3, CNCI, 2, CAS; 6, MCZC), MCZ-ENT00726981, MCZ-ENT00726982, MCZ-ENT00726985 ( E. borealis PARATYPE), MCZ-ENT00727085, MCZ-ENT00727188, MCZ-ENT00727189; Edmonton, 10.v.1916, A.C. Davis (4, CNCI) ; 24 mi. S of Lloydminster, 30.v.1963, Forest Insect Survey, on Populus tremuloides (1, CNCI) ; McMurray, 4.vi.1953, Brooks – Wallis (1, CNCI) ; Exshaw, 27.vi.1954, B.F. and J.L. Carr (1, CNCI) ; Manitoba: Aweme, 28.iv.1903 – 19.v.1922, N. Criddle (1, DEBU; 1, CNCI; 1, MCZC), DEBU01089104, MCZ-ENT00726983; “ Manitoba ” (1, RBCM) , RBCM EENT991-111634; Gillam, 11.vi.–17.vii.1950, W.J. Brown and J.F. McAlpine (27, CNCI) ; Onanole, Riding Mountain National Park, 8.vi.1937 – 15.vi.1938, W.J. Brown (15, CNCI) ; Carberry, 9.v.1953, Brooks – Kelton (1, CNCI) ; Makinak (3, CNCI); New Brunswick: Queens Co., Cranberry Lake P.N.A. (46.1125° N, 65.6075° W), 3–25.v.2011, M. Roy and V. Webster, Red Oak forest Lindgren funnel trap (2, NBM) , NBM-070113 – NBM-070115; Belleville Meduxnekeag Valley Nature Preserve (46.1878° N, 67.6705° W), 18.viii.2008, R.P. Webster (1, NBM) , NBM-070105; Northwest Territories (new territorial record): Norman Wells, 9.iv.1949 – 26.v.1953, S.D. Hicks and C.D. Bird (75, CNCI) ; Ontario: Ridgeway, Liebeck (3, MCZC), MCZ-ENT00726946, MCZ-ENT00726947, MCZ-ENT00727187; Prince Edward Co., 28.v.1919 – 1.vi.1923, J.F. Brimley (11, CNCI); Rainy River District, 14.v.1924, J.F. Brimley (6, CNCI) ; Emo, 21.vi.1924, J.F. Brimley (2, CNCI) ; Constance Bay, 6.viii.1959, H. Howden, on sweetfern (1, CNCI) ; Constance Bay, 17.iv.1968 – 29.iv.1998, Henry and Anne Howden (60, CMNC) ; Constance Bay, 1970, S. Peck (1, CNCI) ; Constance Bay, 21.iv.1977, M. Sanborne (2, CMNC) ; Carleton Co., Constance Bay, 26.iv.1935, W.J. Brown (5, CNCI) ; Constance Bay, 19.v.1933, W.J. Brown (22, CNCI) ; Prince Edward Co., 14.viii.1964, J.F. Brimley (4, UAIC), UAIC1072869; Ottawa, 22.iv.1976, M. Sanborne, on willow (2, CMNC) ; Ottawa, 23.v.1965, A.T. Howden (1, CMNC) ; Ottawa, 6.v.1930, W.J. Brown (3, CNCI) ; Ottawa, 17.v.1961, Kelton (1, CNCI) ; Mer Bleue, 13.v.1932 – 10.v.1933, W.J. Brown (38, CNCI) ; Kenora District, Minnitaki, 12.v.1960, F.I.S., on female Populus tremuloides catkins (2, CNCI) ; Hartington, Eel Lake, South Frontenac (44.563° N, 76.5522° W), 6.ix.2017, J. deWaard (1, CBG) , BIOUG35446 - H11, ELPCH211–17; Hartington, Eel Lake, South Frontenac (44.5645° N, 76.5516° W), 25.vii–8.viii.2017, G. Blagoev (1, CBG) , BIOUG35442 - G10, ELPCG9470-17; Quebec: Cantley, 45.591726° N, 75.784856° W, 17.v.2020, J.H. Lewis, (1, CMNC) , LEW_69, OP215778 (GenBank CO 1 accession number), OP218943 (GenBank ITS 2 accession number); near chemin Holmes (Cantley), 45.598843° N, 75.798396° W, on female Salix catkins, J.H. Lewis, (1, CMNC) , LEW_71, OP215779 (GenBank CO 1 accession number), OP218944 (GenBank ITS 2 accession number); Cantley, 45.59468° N, 75.794389° W, 24.iv.2021, J.H. Lewis, beaten of Salix sp., (1, CMNC), LEW_135, OP215784 (GenBank CO 1 accession number), OP218954 (GenBank ITS 2 accession number); Rigaud, Liebeck Collection (1, MCZC) , MCZ-ENT00726952; Gatineau, near Boucher Forest (45.427673° N, 75.823561° W), 2.v.2021, J.H. Lewis, beaten from Populus tremuloides (2, CMN) , LEW_118, OP215783 (GenBank CO 1 accession number); Gatineau, Mont Cascades (45.588793° N, 75.845822° W), 13.v.2021, J.H. Lewis, beaten from Salix (1, CMNC) , LEW_90, OP215780 (GenBank CO 1 accession number), OP218948 (GenBank ITS 2 accession number); Estrie Region, Scotstown, 14–21.v.2012, C. Levesque (2, CNCI) ; Kirks Ferry, 25.v.1950, B.P. Beirne, in light trap (1, CNCI) ; Laniel, 14.viii.1982, W.J. Brown (1, CNCI) ; Gatineau Park, Harrington Lake, 31.v.1954, R. McCondochie (1, CNCI) ; Cartier, Mount Jacques, 9.vii.1954, W.J. Brown (1, CNCI) ; Kazubazua, 25.v.1933, W.J. Brown (8, CNCI) ; Seven Islands, 8.vi.1929, W.J. Brown (6, CNCI) ; Saskatchewan (new provincial record): Prince Albert, 3.vii.1954, Brooks – Wallis (1, CNCI) ; Fish Creek, 20.v.1928, K.M. King (3, CNCI) ; Elbow, 23.vi.1954, Brooks – Wallis (1, CNCI) ; FINLAND: Espoo, Vanttila (60.1829° N, 24.6157° E), 6.viii.2012, E. Helve (1, ZMUO) , ZMUO.004794, KJ962308; Kuusamo, Aroniemi (66.2741° N, 29.7033° E), 14.vi.2014, Mikko Pentinsaari (2, ZMUO) , ZMUO.016797, ZMUO.016798, COLFH362-14; COLFH363-14; GERMANY: Laucha an der Unstrut, C. Schenkling (4, CNCI); Bavaria, Holzkirchen, 19.v.1957 – 8.v.1959, K. Gaigl (3, ZSM) ; Bavaria, Dietramszell, 26.iv.1959, K. Gaigl (1, ZSM) ; POLAND: Owl Mountains (Eulengeb) (2, MIZ) , COL000397; Sudety-G, Stolowe, Zacisze, 15.v.1947, M. Wegrzecki (1, MIZ) , COL000398; Sierakow, 15.viii.1950, B. Burakowski (1, MIZ) , COL000399; Schlesien (1, MIZ), COL000401; RUSSIA: 2011, B. Korotyeav (3, CMNC); SLOVAKIA: Čačín, Bystrická vrchovina hills (48° 40.3' N, 19° 15.4' E), 6.vii.2020, Michael Košťál (2, CMNC) , LEW_95, OP215781 (GenBank CO 1 accession number), OP218950 ( GenBank ITS 2 accession number), LEW_97, OP215782 (GenBank CO 1 accession number), OP218951 (GenBank CO 1 accession number) . UNITED STATES OF AMERICA: Alaska: Moose Creek (64.71119° N, 147.10143° W), 26.v.–9.vi.2015, J. Hagelin, spruce, meadow, hanging Malaise (1, UAM) , UAM100387793; Connecticut: Cornwall, 14.vii.1925, C.A. Frost (1, MCZC) , MCZ-ENT00727088; Indiana: Morgan Co., 14.vii–7.viii.2012, K.E. Schnepp, Malaise Trap (1, CMNC), LEW_104, OP218952 (GenBank ITS 2 accession number); Elkhart, Liebeck (3, MCZC), MCZ-ENT00726950, MCZ-ENT00726951, MCZ-ENT00700727186; Maine: Hampdan Highlands, 12.viii.1929, H.C. Fall Collection (1, MCZC) , MCZ_ENT00727182; Cumberland Co., 7.v.1916, C.A. Frost Collection (2, MCZC), MCZ-ENT00726970, MCZ-ENT00727073; Massachusetts: Middlesex Co., Natick, 15.vii.1928, C.A. Frost (1, CNCI) ; Middlesex Co., Natick, 15.vii.1928 – 18.iv.1952, C.A. Frost, Salix catkins (1, CMNC; 3, MCZC), MCZ-ENT00726971 – MCZ-ENT00726973; Michigan: Cheboygan Co., Cheboygan, 12.v.1921, S. Moore (1, UAIC) , UAIC1072870; “Mich.”, F.C. Bowditch Collection (1, MCZC), MCZ-ENT00726959; Missouri: St. Louis, 26.iii.1908 – 12.iv.1908, G.W. Bock (3, MCZC) , MCZ-ENT00726949, MCZ-ENT00726977 – MCZ-ENT00726980; New Hampshire: Three Mile Island, 9.vi.1907 – 8.v.1927, Frederick Blanchard (3, MCZC) , MCZ-ENT00727091, MCZ-ENT00727098, MCZ-ENT00727099; Mount Washington, 11.v.1910 – 16.vii.1930, H.C. Fall Collection, Liebeck Collection, F.C. Bowditch Collection and F. Waldo Dodge Collection (11, MCZC) , MCZ-ENT00726944, MCZ-ENT00726948, MCZ-ENT00727080 – MCZ-ENT00727084, MCZ-ENT00727095, MCZ-ENT00726996, MCZ-ENT00727177, MCZ-ENT00727185; Mount Washington, 1874, Frederick Blanchard Collection (3, MCZC), MCZ-ENT00726974 – MCZ-ENT00726976; Farmington, 3.viii.1893 – 30.vii.1900, H.C. Fall Collection (2, MCZC) , MCZ-ENT00700727183, MCZ-ENT00727184; “N.H.”, F.C. Bowditch Collection (4, MCZC), MCZ-ENT00726955 – MCZ-ENT00726958; New York: “N.Y.”, H.C. Fall (2, MCZC), MCZ-ENT00727180, MCZ-ENT00727181; Oregon: Burgess, A. Fenyes (1, CAS); Pennsylvania: Hazleton, Liebeck Collection (1, MCZC), MCZ-ENT00726984; Allegheny Co., Frederick Blanchard Collection and W.G. Dietz Collection (6, MCZC), MCZ-ENT00726967, MCZ-ENT00727094, MCZ-ENT00727100 – MCZ-ENT00727103; “Pa.”, H.C. Fall Collection, F.C. Bowditch Collection, and W.G. Dietz (6, MCZC), MCZ-ENT00726953, MCZ-ENT00726954, MCZ-ENT00726966, MCZ-ENT00727093, MCZ-ENT00727094, MCZ-ENT00727179.

Diagnosis. Length 2.5–2.9 mm. Cuticular colouration black and/or red in most specimens; rarely entirely pale yellow. Scales on pronotum uniform in colour (usually white) or only with central band of discolouration; lacking discoloured lateral bands. Metaventrite and metepisternum with scales relatively sparse, not obscuring underlying cuticle. Shiny carina running longitudinally along last ventrite of males (at least present as swelling at the middle of ventrite). Apex of aedeagus smoothly rounded (semicircular); not subquadrate or distinctly pointed. Internal sac tuberculate, with distinct sclerotised hook like structure; lacking any protruding basal structures. Distributed across temperate and boreal North America.

Natural history. This species has been collected on Populus tremuloides (Michaux) and Salix bebbiana Sargent in North America, and in Europe from S. capraea Linnaeus, S. viminalis Linnaeus, S. cinerea Linnaeus, and S. aurita Linnaeus (Hoffmann 1954) . Specimens of E. bipunctatus have also been collected in Lindgren funnel traps set in the canopy of P. grandidentata Michaux in New Brunswick (Webster et al. 2016) and in fermenting bait traps (beer/wine and sugars) in Russia (Ruchin et al. 2021).

Distribution in North America. This species is distributed widely across Canada, from New Brunswick to the Northwest Territories, and is also known from several American states, including Alaska and Oregon, and east of the Great Plains.

Remarks. Molecular data indicate some level of differentiation between the European and North American populations of this species (see Figs. 1, 2); however, dissections and examinations of external morphology did not uncover any significant differences in morphology warranting formal naming. The limited CO 1 and ITS 2 data available (Figs. 1, 2) do not suggest North American and European populations mixing (i.e., introduction events), although broader sampling could be conducted to further investigate this possibility. Here, we treat these as the same species. This has implications for “ E. borealis ” (Carr, 1920), which is determined here to represent an all-red male E. bipunctatus on the basis of the following: (1) possessing a conspicuous, shiny carina extending longitudinally over the last ventrite (Fig. 3A), (2) with the metepisternum and lateral half of the metaventrite not densely punctate or scaled, and with the cuticle in those regions mostly visible (Fig. 3D), and (3) dissections of similar, all-red specimens from the Canadian Prairies are all E. bipunctatus . For this reason, E. borealis is hereafter treated as a junior synonym of E. bipunctatus . Label data for the holotype of E. borealis (examined) are as follows: CANADA, Alberta, Edmonton, 24.v.1919, F.S. Carr [Edmonton Alta 24.v.1919 F.S. Carr]. Type Elleschus borealis, No. 419. CNC 379701.